Porifera, Demospongiae)

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Porifera, Demospongiae) Bijdragen tot de Dierkunde, 52 (2): 82-102 — 1982 Ecology and distribution of two sympatric, closely related sponge species, Halichondria panicea (Pallas, 1766) and H. bowerbanki Burton, 1930 (Porifera, Demospongiae), with remarks on their speciation by A.D. Vethaak R.J.A. CronieI & R.W.M. van Soest Institute of Taxonomie Zoology, University of Amsterdam, P.O. Box 20125, 1000 HC Amsterdam, The Netherlands Abstract INTRODUCTION The occurrence of two or more closely North Atlantic coastal waters harbour two closely related sympatric the viz. sympatric sponges belonging to genus Halichondria, related species has inspired the development of H. panicea and H. bowerbanki. From the study of their speciation theories in which ecological niche dif- littoral and sublittoral microdistribution and ecology in the ferentiation a role 1974: Oosterschelde = Eastern S.W. of The area ( Scheldt, part plays major (e.g. Ross, Netherlands) it is concluded that both similar occupy largely 101-106). Such a presumed mode of speciation found better niches, although it was that H. panicea is does not necessitate geographical isolation of to endure to air since it equipped exposure occurs up to a vicariance or higher intertidal level than H. bowerbanki, while the latter populations (through dispersal), is better withstand equipped to siltation. Other environmental which is normally considered the most parsi- factors as depth, light, current velocity and salinity affect the monious of way speciation. Sympatric speciation distribution and behaviour of both species similarly. From is made feasible in certain these observations can be quite parasitic insects, it inferred, that speciation from a parent species cannot have been sympatrically through but convincing unequivocal examples in other ecological segregation of individuals. The possibilities of animal In marine groups or habitats are rare. the isolation of geographical of populations a parent species on sides of environment of both the Atlantic Ocean during past glacial periods suggestions sympatric speciation have resulting in an originally American species ( H . bowerbanki) been made by Sara (1956, 1957) when and an originally European species ( H. panicea) are dis- trying to explain the sympatric occurrence of the cussed. calcareous sponges Leucosolenia complicata (Mon- tagu, 1818), L. variabilis Haeckel, 1870 and L. Résumé botryoides (Ellis & Solander, 1786) in western Europe, and by McGowan (1963) when trying Les eaux côtières de l’Atlantique Nord hébergent deux fort de to the occurrence of the espèces sympatriques apparentées Spongiaires apparte- explain sympatric pelagic nant au genre Halichondria, à savoir H. panicea et H. bower- molluscs Limacina helicina (Phipps, 1774) banki. L’étude de leur microdistribution et de leur écologie varieties A and in the dans le littoral B, North Pacific. et le sublittoral de la zone de l’Oosterschelde ( = des A indication for l’Escaut oriental, partie S.O. Pays-Bas) permet de strong a possible sympatric conclure les deux des niches en général simi- que occupent is the niche differentiation such speciation a group laires; on a cependant trouvé que H. panicea est mieux of related should exhibit. This à à closely species adaptée l’exposition l’air, puisqu’elle peut être présente niveau jusqu’à un intercotidal supérieur à celui atteint indication is, still not conclusive because par however, H. bowerbanki, tandis cette dernière est que espèce mieux niche differentiation have may just as well résister préparée pour à l’envasement. D’autres facteurs du occurred after the event instead of comme la speciation being milieu, profondeur, la lumière, la vitesse du cou- la salinité, affectent de the cause of it. At without rant, manière similaire la distribution any rate, such a clear et le comportement des deux espèces. déduire On peut de ces niche differentiation a theory of sympatric specia- observations que, dans ce cas, la spéciation à partir d’une tion (difficult to in would espèce parentale n’a être enough explain itself) pas pu sympatrique, par ségrégation des écologique individus. On discute des possibilités d’isola- be unwarranted. tion géographique de populations d’une espèce parentale des The alternative theory, speciation through deux côtés de l’Atlantique au cours des périodes glaciaires, has been used to comme résultat une geographical isolation, explain ayant espèce d’origine américaine (H. bowerbanki) et une autre marine but d’origine européenne (H. panicea) . speciation events, not very widely - 1982 83 BIJDRAGEN TOT DE DIERKUNDE, 52 (2) it rather of the was because involves a intimate knowledge Sampling sponge populations per- of past geological and climatological events and formed by hand, hand-operated grabs, snorkeling, the SCUBA and data on the dispersal capabilities of species diving dredging. concerned. In tropical parts of the oceans these The following preserved material from the collections of events are known to a modest we can degree: the Zoölogisch Museum, University of Amsterdam (ZMA), Historie at Leiden vicariant distributions of the van Natuurlijke explain many closely Rijksmuseum (RMNH), the British Museum (Natural History), London related benthic marine taxa by the former existence (BMNH) and the Zoologisk Museum, University of Copen- Sea and its of the Tethys subsequent disruption the hagen (ZMUC) has been used in present study: because of the openings of the Atlantic Ocean Halichondria for panicea: (e.g. Ekman, 1953; Briggs, 1974; sponges: ZMA (The Netherlands): Por. 2985 through 3035; 3063; Wiedenmayer, 1977). In the pelagic environment 3069; 3070; 3077; 3079; 3086; 3093; 3094; 3101; such vicariance theories have been formulated by 3103; 3115; 3123; 3128; 3131 through 3135; 3152; Brinton (1962), Van Soest (1975), and Pierrot 3157; 3161 through 3169; 3179; 3182 through 3185; 3220; 3224; 3226; 3229; 3234; 3246; 3466; 3471; & Van der Spoel (1979). In most individual 3973; 4083; 4105 through 4107; 4109; 4111 through is lack of sufficient cases, however, there always a 4115; 4162; 4175; 4202; 4207; 4209; 4216; 4221; data the the in- 4223; 4227; 4329; 4330; 4332; 4336; 4449 through on past distribution of species 4451; 4478; 4574; 4575. volved and the geological events that should have ZMA (Ireland): Por. 4426; 4689; 4690. led the of of the to segregation populations ZMA (New Zealand; donated by P. Bergquist, Auckland ancestral species. University): Por. 4553. ZMA (France): Por. 2485; 2496; 2498. In will the following we try to reconstruct the RMNH (The Netherlands): Nos. 423 through 432; 1075 scenario of two related evolutionary closely sponge through 1081; 1082 through 1085; 1086; 1087 species, viz. Halichondria panicea (Pallas, 1766) through 1091; 1156 through 1159; 1162; 1164; 1181 and through 1183; 1191; 1193; 1207; 1209; H. bowerbanki Burton, 1930, which occur 1210 through 1213; 1218; 1219; 1222; 1224; in most of their sympatrically geographic ranges. 1230; 1231; 1255; 1263 through 1265; 1275; Because literature data niche 22 unnumbered. suggest a differen- 1280; samples, BMNH material, labeled Halichondria panicea: Sealark tiation in these species it will be attempted to Kxped., Indian Ocean; Lambert Bay, S. Africa; show at that the of both length, ecology species Pt. Noloth, S. Africa (1938.3.26.72); Manihine precludes the assumption of sympatric speciation. Exped., Gulf of Aqaba; Challenger Exped., Ker- guelen, Indian Ocean (1887.6.2.216); Challenger will Subsequently an alternative way of speciation Exped., Japan; Mergui Archipel, Birma; Cape be discussed. The value of our results predictive Adare, Antarctica; Japanese Sea; Greenland; Es- Harbour, is unknown, although several similar cases of quimault Vancouver, W. Canada; S. Sa- chalin, U.S.S.R. (1931.6.10.10); Inverness Wood- closely related, sympatric, marine benthic species cock's Landing, Estuary of the Sheena River, British in other animal known exist. pairs groups are to Columbia, W. Canada (1890.9.2.1); Swedish Ant- arctic Exped., Falkland Isis. (1933.3.17.19; no ACKNOWLEDGEMENTS sponge). ZMUC Atlantide Exped., Sta. 146 (9°24'N 14°45'W), The authors are very much indebted to Mr. Jan Vermeulen for 50-51 m. his excellent assistance in the field. Furthermore, we are grateful to Mr. J. Zaagman for retouching the photographic Halichondria bowerbanki: illustrations and for assembling the plates; Mr. H. Mittelberg made the ZMA Por. 3146 frame box apparatus. Miss S. M. Stone (British (The Netherlands): 2681; 3138; through Museum (Natural History)), Dr. P. R. Bergquist (Auckland, 3148; 3472; 4087 through 4090; 4099; 4101; 4103; New and 4116 through 4118; Zealand) Dr. E. Rasmussen (Copenhagen) are 4122 through 4125; 4132; 4133; thanked for putting material at our disposal. 4199; 4203; 4219; 4220; 4224; 4328; 4331; 4334; 4337; 4472; 4574; 4575. ZMA (Ireland): Por. 4297; 4421; 4547 MATERIALS AND METHODS ZMA (France, as H. coali ta) : Por. 2490 through 2495; fieldwork The part of this study was carried out 2499. by visits to 17 localities in the making regular BMNH: Plymouth; Naples (Italy). BMNH-material labeled Oosterschei de “Halichondria lacunosa”: (Eastern Scheldt) area during a Port Chevalier Reef, New Zealand (1938.8.24.12). BMNH-material labeled three-year period from November 1977 till “Halichondria caduca”: Challenger Exped., S.W. December 1980. Patagonia. 84 A. D. VETHAAK ET AL. - SYMPATRIC HALICHONDRIA SPECIES the deter- 2 from the basal Reproductive periods of sponges were 0.5 cm were cut marginal zone mined the and of collected in 5, during years 1978, 1979 1980,
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